山东省12个主栽小麦品种(系)抗叶锈性分析

本研究旨在明确山东省12个小麦主栽品种(系)抗叶锈性及抗叶锈基因,为小麦品种推广与合理布局、叶锈病防治及抗病育种提供依据。利用2015年采自山东省的5个小麦叶锈菌流行小种的混合小种对这些材料进行苗期抗性鉴定,然后选用15个小麦叶锈菌生理小种对这些品种(系)进行苗期基因推导,并利用与24个小麦抗叶锈基因紧密连锁(或共分离)的30个分子标记对其进行抗叶锈基因分子检测。结果显示,山东省12个主栽小麦品种(系)苗期对该省2015年的5个小麦叶锈菌混合流行小种均表现高度感病。通过基因推导与分子检测发现,济南17含有Lr16,矮抗58和山农20含有Lr26,其余济麦系列、烟农系列、良星系列等9个品种(系)均未检测到所供试标记片段。此外,本研究还对山东省3个非主栽品种进行了检测,结果发现,中麦175含有抗叶锈基因Lr1和Lr37,含有成株抗性基因;皖麦38只检测到Lr26,济麦20未检测到所供试标记片段。综合以上结果,山东省主栽小麦品种(系)所含抗叶锈基因丰富度较低,尤其不含有对我国小麦叶锈菌流行小种有效的抗锈基因,应该引起高度重视,今后育种工作应注重引入其他抗叶锈基因,提高抗叶锈性。     

优异大麦种质资源的抗性鉴定和评价

1998～1999年,将1997年筛选出的191个农艺性状表现优良的大麦品种进行大麦黄花叶病、大麦白粉病、大麦赤霉病、大麦条纹病及耐盐等抗性的鉴定评价,结果表明,抗及高抗大麦黄花叶病品种60份,抗大麦赤霉病品种59份,高抗大麦条纹病品种89份,抗大麦白粉病品种19份,芽期耐盐品种17份,苗期耐盐品种2份.     

优异大麦种质资源的抗性鉴定和评价

1998～1999年,将1997年筛选出的191个农艺性状表现优良的大麦品种进行大麦黄花叶病、大麦白粉病、大麦赤霉病、大麦条纹病及耐盐等抗性的鉴定评价,结果表明,抗及高抗大麦黄花叶病品种60份,抗大麦赤霉病品种59份,高抗大麦条纹病品种89份,抗大麦白粉病品种19份,芽期耐盐品种17份,苗期耐盐品种2份。     

河南省16个主栽小麦品种抗叶锈基因分析

为了明确河南省小麦品种的抗叶锈性及抗叶锈基因的分布,为小麦品种推广与合理布局、叶锈病防治及抗病育种提供依据,本研究利用2015年采自河南省的5个小麦叶锈菌流行小种混合菌株,对近几年河南省16个主栽小麦品种进行了苗期抗性鉴定,然后选用12个小麦叶锈菌生理小种对这些品种进行苗期基因推导,同时利用与24个小麦抗叶锈基因紧密连锁(或共分离)的30个分子标记对该16个品种进行了抗叶锈基因分子检测。结果显示,供试品种苗期对小麦叶锈菌混合流行小种均表现高度感病;基因推导与分子检测结果表明,供试品种可能含有Lr1、Lr16、Lr26和Lr30这4个抗叶锈基因,其中先麦8号含有Lr1和Lr26;郑麦366和郑麦9023含有Lr1;西农979和怀川916含有Lr16;中麦895、偃展4110、郑麦7698、平安8号、众麦1号、周麦16、衡观35和矮抗58含有Lr26;周麦22中含有Lr26,还可能含有Lr1和Lr30;豫麦49-198和洛麦23可能含有本研究中检测以外的其他抗叶锈基因。因此,河南省主栽小麦品种的抗叶锈基因丰富度较低,今后育种工作应注重引入其他抗叶锈性基因,提高抗叶锈性,有效控制小麦叶锈病。     

小麦品种梭条花叶病抗性基因遗传分析及分子标记筛选

选用3个抗梭条花叶病的小麦品种“仪宁小麦”、“徐87633”和“西风”为抗病亲本、以感病品种“镇9523”为感病亲本配制了3个杂交组合,对4个亲本及其杂种后代(F1及F2代)单株的田间抗病鉴定表明,3个抗病亲本的抗性均由核基因控制,为显性遗传方式。“仪宁小麦”和“西风”的抗性受两对表现互补效应的显性基因控制;“徐87633”的抗性受一对显性基因控制。选取涉及小麦21条染色体上的266对SSR引物在“仪宁小麦”和“镇9523”间进行筛选,其中108对引物在两亲本间表现多态。根据“仪宁小麦”ד镇9523”F2代单株的田间抗病鉴定结果,采用BSA的方法,将已初筛的108对引物在抗、感池间进行扩增,发现引物Xgwm261在抗、感池间表现多态,表明该引物与“仪宁小麦”的抗病基因连锁,并将该抗病位点初步定位于2DS上。用该标记对F2代224个单株进行PCR扩增,根据扩增结果,采用Mapmaker3.0软件计算遗传距离,结果显示,该标记与抗病位点间的遗传距离为22.9cM。     

小麦品种莱州137抗叶锈性鉴定及基因检测

摘要：小麦叶锈病是小麦生产中的重要病害,培育持久抗叶锈性品种可以有效、经济地控制该病害。本文通过基因推导、分子检测结合系谱分析成株抗性鉴定对小麦重要生产品种中抗病基因进行分析,从而确定小麦品种中所携带的抗病基因。本试验用20个不同毒力的叶锈菌菌系、36个已知抗叶锈基因载体品种以及感病对照品种郑州5389对供试品种莱州137进行苗期抗叶锈病基因推导分析,并分别用12个与抗叶锈病基因连锁的分子标记进行目的基因的分子检测,同时利用系谱分析法来验证莱州137所携带的已知抗叶锈病基因;在2014-2015年度和2015-2016年度将莱州137、慢锈性对照品种SAAR和感病对照品种郑州5389种植于河北农业大学小麦试验田和河南周口黄泛区农场试验田,用田间混合生理小种（FHRT、THTT、THJT）接种进行成株期抗病性鉴定。结果表明,通过苗期基因推导分析,莱州137对小种FGBQ、PGJQ、TGTT、THSM、PHGM、PHST、FHJS、FHGQ、FNTQ、PRSQ和KHGQ表现抗病,而Lr26对FGBQ、PGJQ和TGTT高抗,Lr10和Lr14b分别对小种THSM、PHGM和PHST、PHGM、THTT表现高抗,同时系谱分析和分子检测也验证已知抗叶锈基因Lr26和Lr10,因此在供试品种莱州137中鉴定出Lr26、Lr10、Lr14b以及未知的抗叶锈病基因;根据2年2点的田间抗叶锈病鉴定,莱州137表现出成株抗性特点,且经分子标记检测该品种中未含有Lr34和Lr46,故小麦品种莱州137中含有未知的成株抗叶锈性基因,可作为新的小麦叶锈病抗源加以利用。     

182份黄淮海麦区小麦品种（系）苗期抗叶锈病基因分析

利用KHST、FHKT和FHJT②3个小麦叶锈菌混合菌株对182份小麦品种(系)进行苗期抗叶锈鉴定,对筛选出的抗性品种利用15个小麦叶锈菌生理小种进行基因推导,结合与20个抗叶锈基因连锁的25个分子标记进行抗叶锈基因分析。182份小麦品种(系)中,14个品种(泰科麦5303、驻麦305、豫圣麦118、存麦18号、轩麦6号、农丰川、丹麦118、郑麦103、郑麦119、赛德麦5号、郑麦369、许科918、豫麦668和AF116-120)表现抗性,其余品种(系)均表现高感;基因推导结果显示,驻麦305、存麦18号、农丰川、郑麦119、赛德麦5号、郑麦369、许科918含有抗叶锈基因Lr33+34;郑麦103含有抗叶锈基因Lr10和Lr33+34;AF116-120含有抗叶锈基因Lr10、Lr16、Lr20和Lr33+34;泰科麦5303、豫圣麦118、丹麦118和豫麦668可能含有其他抗叶锈基因;分子检测结果显示,农川丰、轩麦6号、郑麦103和许科918含有抗叶锈基因Lr1和Lr26;泰科麦5303、豫圣麦118、郑麦119和郑麦369含有抗叶锈基因Lr1;驻麦305、存麦18号和豫麦668含有抗叶锈基因Lr26;AF116-120含有抗叶锈基因Lr1和Lr2c;丹麦118含有抗叶锈基因Lr26和Lr37。所检测小麦品种含抗叶锈基因丰富度低,缺乏有效抗叶锈基因。182份黄淮海麦区小麦品种对小麦叶锈菌的抗病性及抗性品种中抗性基因组成的分析,可以为该地区小麦品种推广、合理布局及叶锈病防治与抗病育种提供科学依据。     

定向改良超级早稻中早39稻瘟病抗性

稻瘟病是水稻生产中最具毁灭性的病害之一,稻瘟病菌生理小种变异快,抗性品种推广3～5年后抗性衰退风险较高,改良及培育抗稻瘟病菌新致病小种的品种是防治该病害最经济有效的方法。本研究以携带稻瘟病抗性基因Pi25的材料R6为供体亲本,超级早稻中早39为受体亲本和轮回亲本,通过杂交、回交和自交以及分子标记辅助选择技术,并结合室内喷雾接种、田间注射接种叶瘟以及病圃自然发病区穗颈瘟鉴定。综合考量抗性表型、基因型以及农艺性状,筛选出4个携带Pi25基因,叶瘟及穗颈瘟抗性较中早39提高,且农艺性状良好的株系FY82、FY90、FY125、FY137,同时对育成品系的系谱组成和抗性相关位点进行研究,初步阐明了育成品系较轮回亲本抗性水平提高的遗传基础。     

小麦抗条锈病基因YrCN19的诊断检测和遗传分析

用小麦条锈病抗性基因YrCN19的诊断标记Xgwm410对19个小麦品种或品系进行PCR筛选,其中川农19、新抗5号、爱民5号和爱民6号扩增出与条锈病抗性基因YrCN19共分离的特征片断,其大小为391个碱基,而在其他的小麦品种或品系中未能检测到该片断。系谱分析和抗性鉴定结果表明川农19、新抗5号、爱民5号和爱民6号含有小麦条锈病基因YrCN19。抗性遗传分析发现小麦条锈病抗性在川农19,新抗5号和爱民5号中的遗传符合单个显性基因的遗传规律(3抗:1感);杂交组合烟辐188/爱民6号的抗性遗传也符合单个显性基因的遗传规律,而另外一些杂交组合(如R25/爱民6号,鲁955159/爱民6号和苏3110/爱民6号)中的抗性分离则符合两对基因互补的遗传规律(9抗:7感)。本研究揭示了小麦条锈病抗性基因YrCN19在不同遗传背景和杂交组合的抗性表达和分离有差异,从而加速YrCN19在小麦抗条锈育种中的开发与利用。     

小麦品种莱州137抗叶锈病鉴定及基因检测

叶锈病是小麦生产中的重要病害,培育持久抗性品种可以有效控制该病害。本研究以抗病品种莱州137、感病对照品种郑州5389、慢锈性品种SAAR以及36个已知抗叶锈病基因的载体品种为材料,在苗期和成株期进行了2年2点的接种鉴定,通过系谱分析、基因推导和12个与抗叶锈病基因连锁的分子标记检测,发现莱州137携带Lr26、Lr10、Lr14b以及其他未知抗叶锈病基因,且表现成株抗性的特点,说明其可能含有未知的成株抗性基因,可作为新的小麦抗叶锈病抗源加以利用。     

Mapping quantitative trait loci associated with spot blotch and net blotch resistance in a doubled-haploid barley population

Spot blotch and net blotch are important foliar barley (Hordeum vulgare L.) diseases in Canada and elsewhere. These diseases result in significant yield reduction and, more importantly, loss of grain quality, downgrading barley from malt to feed. Combining resistance to these diseases is a breeding priority but is a significant challenge using conventional breeding methodology. In the present investigation, an evaluation of the inheritance of resistance to spot and net blotch was conducted in a doubled-haploid barley population from the cross CDC Bold (susceptible)?×?TR251 (resistant). The population was screened at the seedling stage in the Phytotron and at the adult-plant stage in the field for several years. Chi-squared analysis indicated one- to four-gene segregation depending on disease, isolate, plant development stage, location and year. A major seedling and adult-plant resistance quantitative trait locus (QTL), designated QRpt6, was re-confirmed for net-form net blotch resistance, explaining 32?C61% of phenotypic variation in different experiments. Additional QTL for seedling and adult-plant resistance to net blotch were identified. For spot blotch resistance, a major seedling resistance QTL (QRcss1) was detected on chromosome 1H for isolate WRS1909, explaining 79% of the phenotypic variation. A highly significant QTL on 3H (QRcs3) was identified for seedling resistance to isolate WRS1908 and adult-plant resistance at Brandon, MB, Canada in 2008. The identification of QTL at only one location or from 1?year suggests spot blotch resistance is complex and highly influenced by the environment. Efforts are being made to combine spot and net blotch resistance in elite barley lines using molecular marker-assisted selection.     

Mapping quantitative trait loci associated with barley net blotch resistance

Net blotch of barley, caused by Pyrenophora teres Drechs., is an important foliar disease worldwide. Deployment of resistant cultivars is the most economic and eco-friendly control method. This report describes mapping of quantitative trait loci (QTL) associated with net blotch resistance in a doubled-haploid (DH) barley population using diversity arrays technology (DArT) markers. One hundred and fifty DH lines from the cross CDC Dolly (susceptible)/TR251 (resistant) were screened as seedlings in controlled environments with net-form net blotch (NFNB) isolates WRS858 and WRS1607 and spot-form net blotch (SFNB) isolate WRS857. The population was also screened at the adult-plant stage for NFNB resistance in the field in 2005 and 2006. A high-density genetic linkage map of 90 DH lines was constructed using 457 DArT and 11 SSR markers. A major NFNB seedling resistance QTL, designated QRpt6, was mapped to chromosome 6H for isolates WRS858 and WRS1607. QRpt6 was associated with adult-plant resistance in the 2005 and 2006 field trials. Additional QTL for NFNB seedling resistance to the more virulent isolate WRS858 were identified on chromosomes 2H, 4H, and 5H. A seedling resistance QTL (QRpts4) for the SFNB isolate WRS857 was detected on chromosome 4H as was a significant QTL (QRpt7) on chromosome 7H. Three QTL (QRpt6, QRpts4, QRpt7) were associated with resistance to both net blotch forms and lines with one or more of these demonstrated improved resistance. Simple sequence repeat (SSR) markers tightly linked to QRpt6 and QRpts4 were identified and validated in an unrelated barley population. The major 6H QTL, QRpt6, may provide adequate NFNB field resistance in western Canada and could be routinely selected for using molecular markers in a practical breeding program.     

Mapping spot blotch resistance genes in four barley populations

Bipolaris sorokiniana (teleomorph: Cochliobolus sativus) is the fungal pathogen responsible for spot blotch in barley (Hordeum vulgare L.) and occurs worldwide in warmer, humid growing conditions. Current Australian barley varieties are largely susceptible to this disease and attempts are being made to introduce sources of resistance from North America. In this study we have compared chromosomal locations of spot blotch resistance reactions in four North American two-rowed barley lines; the North Dakota lines ND11231-12 and ND11231-11 and the Canadian lines TR251 and WPG8412-9-2-1. Diversity arrays technology-based PCR, expressed sequence tag and SSR markers have been mapped across four populations derived from crosses between susceptible parental lines and these four resistant parents to determine the location of resistance loci. Quantitative trait loci (QTL) conferring resistance to spot blotch in adult plants (APR) were detected on chromosomes 3HS and 7HS. In contrast, seedling resistance (SLR) was controlled solely by a locus on chromosome 7HS. The phenotypic variance explained by the APR QTL on 3HS was between 16 and 25% and the phenotypic variance explained by the 7HS APR QTL was between 8 and 42% across the four populations. The SLR QTL on 7HS explained between 52 and 64% of the phenotypic variance. An examination of the pedigrees of these resistance sources supports the common identity of resistance in these lines and indicates that only a limited number of major resistance loci are available in current two-rowed germplasm.     

Association mapping of spot blotch resistance in wild barley

Spot blotch, caused by Cochliobolus sativus, is an important foliar disease of barley. The disease has been controlled for over 40 years through the deployment of cultivars with durable resistance derived from the line NDB112. Pathotypes of C. sativus with virulence for the NDB112 resistance have been detected in Canada; thus, many commercial cultivars are vulnerable to spot blotch epidemics. To increase the diversity of spot blotch resistance in cultivated barley, we evaluated 318 diverse wild barley accessions comprising the Wild Barley Diversity Collection (WBDC) for reaction to C. sativus at the seedling stage and utilized an association mapping (AM) approach to identify and map resistance loci. A high frequency of resistance was found in the WBDC as 95% (302/318) of the accessions exhibited low infection responses. The WBDC was genotyped with 558 Diversity Array Technology (DArT^®) and 2,878 single nucleotide polymorphism (SNP) markers and subjected to structure analysis before running the AM procedure. Thirteen QTL for spot blotch resistance were identified with DArT and SNP markers. These QTL were found on chromosomes 1H, 2H, 3H, 5H, and 7H and explained from 2.3 to 3.9% of the phenotypic variance. Nearly half of the identified QTL mapped to chromosome bins where spot blotch resistance loci were previously reported, offering some validation for the AM approach. The other QTL mapped to unique genomic regions and may represent new spot blotch resistance loci. This study demonstrates that AM is an effective technique for identifying and mapping QTL for disease resistance in a wild crop progenitor.     

Genome-wide association mapping reveals genetic architecture of durable spot blotch resistance in US barley breeding germplasm

Spot blotch, an economically important disease of both barley and wheat, is caused by Cochliobolus sativus (anamorph: Bipolaris sorokiniana). The disease has been reported in many regions of the world, but is particularly severe on barley in the Upper Midwest region of the USA and adjacent areas of Canada. For over 50 years, spot blotch has been effectively controlled through the deployment of durable resistance in six-rowed malting cultivars. To characterize loci conferring spot blotch resistance in US barley germplasm, we employed an association mapping approach using 3,840 breeding lines and cultivars. Three quantitative trait loci (QTL), Rcs-qtl-1H-11_10764, Rcs-qtl-3H-11_10565 and Rcs-qtl-7H-11_20162, were found to confer both seedling and adult plant resistance. Together, these three QTL comprise the Midwest Six-rowed Durable Resistant Haplotype (MSDRH), which is present in all Midwest six-rowed cultivars released since the 1960s. Each QTL alone only partially reduced disease levels, but combining all three together reduced the seedling infection response and adult plant disease severity by 47 and 83 %, respectively. The identified MSDRH will be valuable for marker-assisted selection of breeding lines to deploy spot blotch resistance and can also be incorporated into genomic selection as one of the disease resistance traits.     

Gene action for adult-plant resistance to powdery mildew in wheat.

Gene action for adult-plant resistance to powdery mildew was studied using generation mean analyses of parents and of F1, F2, and backcross populations derived from a diallel cross of one susceptible and three adult-plant resistant wheat cultivars. Joint scaling tests showed that an additive-dominance model was sufficient to explain the variability in the expression of adult-plant resistance in one cross, while digenic epistasis was involved in the other five crosses. Additive gene effects were predominant; however, dominance was significant in four crosses, additive x additive interaction was significant in three crosses, additive x dominance interaction was significant in three crosses, and dominance x dominance interaction was significant in one cross. Therefore, selection for adult-plant resistance would likely be most effective in advanced generations derived from crosses among the adult-plant resistant cultivars Redcoat, Houser, and Massey.     

Genetics of seedling and adult plant resistance to net blotch (Pyrenophora teres f. teres) and spot blotch (Cochliobolus sativus) in barley

Net blotch (caused by Pyrenophora teres f. teres) and spot blotch (Cochliobolus sativus) are important foliar diseases of barley in the midwestern region of the USA. To determine the number and chromosomal location of Mendelian and quantitative trait loci (QTL) controlling resistance to these diseases, a doubled haploid population (Steptoe/Morex) was evaluated to the pathogens at the seedling stage in the greenhouse and at the adult plant stage in the field. Alleles at two or three unlinked loci were found to confer resistance to the net blotch pathogen at the seedling stage depending on how progeny exhibiting an intermediate infection response were classified. This result was corroborated in the quantitative analysis of the raw infection response data as 2 major QTL were identified on chromosomes 4 and 6M. A third QTL was also identified on chromosome 6P. Seven QTL were identified for net blotch resistance at the adult plant stage and mapped to chromosomes 1P, 2P, 3P, 3M, 4, 6P, and 7P. The 7 QTL collectively accounted for 67.6% of the phenotypic variance under a multiple QTL model. Resistance to the spot blotch pathogen was conferred by a single gene at the seedling stage. This gene was mapped to the distal region of chromosome 1P on the basis of both qualitative and quantitative data analyses. Two QTL were identified for spot blotch resistance at the adult plant stage: the largest QTL effect mapped to chromosome 5P and the other mapped to chromosome 1P near the seedling resistance locus. Together, the 2 QTL explained 70.1% of the phenotypic variance under a multiple QTL model. On the basis of the chromosomal locations of resistance alleles detected in this study, it should be feasible to combine high levels of resistance to both P. teres f. teres and C. sativus in barley cultivars.     

Progress in Wheat Resistance to Spot Blotch in Bangladesh

Spot blotch, caused by Cochliobolus sativus, is considered one of the most destructive diseases of wheat (Triticum aestivum) in the warm areas of South Asia. Over the past 20 years, wheat breeding efforts in the region have improved spot blotch resistance in susceptible commercial cultivars. This study assessed resistance and spot blotch‐induced yield losses in newly released wheat cultivars developed in Bangladesh since the release of the landmark wheat variety ‘Kanchan’. Replicated field studies were conducted during the 2003 and 2004 wheat seasons at two sites: a farmer's field and a research station in a warm region of Bangladesh where spot blotch has been a serious problem. Spot blotch affected 60% of the crop and caused yield losses of from 2% to 22%. Disease severity and disease‐induced grain yield reductions were less in wheat genotypes developed since 1983, with a corresponding trend towards higher yield in newly developed varieties. The level of resistance to spot blotch in the new cultivars and advanced breeding lines represents considerable progress in breeding for resistance over the past two decades.     

Inheritance and RAPD tagging of multiple genes for resistance to net blotch in barley.

A doubled haploid barley (Hordeum vulgare L.) population that was created from a cross between cultivars 'Léger' and 'CI 9831' was characterized by RAPD (random amplified polymorphic DNA) markers for resistance to isolate WRS857 of Pyrenophora teres Drechs. f. sp. maculata Smedeg., the causal agent of the spot form of net blotch. Resistance, which initially appeared to be conferred by a single gene from the approximate 1:1 (resistant : susceptible) segregation ratio of the doubled-haploid (DH) progeny, was found to be associated with three different genomic regions by RAPD analysis. Of 500 RAPD random primers that were screened against the parents, 195 revealed polymorphic bands, seven showed an association to the resistance in bulks, and these seven markers were mapped to three unlinked genomic regions. Two of these regions, one of which was mapped to chromosome 2, have major resistance genes. The third region has some homology to the chromosome 2 region. This study demonstrates the simultaneous location of markers for more than one gene governing a trait by using RAPD and bulked segregant analysis (BSA).     

Prospects for exploitation of disease resistance from Hordeum chilense in cultivated cereals

Hordeum chilense is a South American wild barley with high potential for cereal breeding given its high crossability with other members of the Triticeae. In the present paper we consider the resistance of H. chilense to several fungal diseases and the prospects for its transference to cultivated cereals. All H. chilense accessions studied are resistant to the barley, wheat and rye brown rusts, the powdery mildews of wheat, barley, rye and oat, to Septoria leaf blotch, common bunt and to loose smuts, which suggests that H. chilense is a non-host of these diseases. There are also lines resistant to wheat and barley yellow rust, stem rust and to Agropyron leaf rust, as well as lines giving moderate levels of resistance to Septoria glume blotch, tan spot and Fusarium head blight. Some H. chilense lines display pre-appressorial avoidance to brown rust. Lines differ in the degree of haustorium formation by rust and mildew fungi they permit, and in the degree to which a hypersensitive response occurs after haustoria are formed. Unfortunately, resistance of H. chilense to rust fungi is not expressed in tritordeum hybrids, nor in chromosome addition lines in wheat. In tritordeum, H. chilense contributes quantitative resistance to wheat powdery mildew, tan spot and loose smut. The resistance to mildew, expressed as a reduced disease severity, is not associated with macroscopically visible necrosis. Hexaploid tritordeums are immune to Septoria leaf blotch and to common bunt although resistance to both is slightly diluted in octoploid tritordeums. Studies with addition lines in wheat indicate that the resistance of H. chilense to powdery mildew, Septoria leaf blotch and common bunt is of broad genetic basis, conferred by genes present on various chromosomes.